In Search of the Sister Group of the True Lice: a Systematic Review of Booklice and Their Relatives, with an Updated Checklist of Liposcelididae (Insecta: Psocodea)

YOSHIZAWA & LIENHARD: Systematics of Liposcelididae In search of the sister group of the true lice: A systematic review of booklice and their relatives, with an updated checklist of Liposcelididae (Insecta: Psocodea) Kazunori Yoshizawa 1,* & Charles Lienhard 2 1 Systematic Entomology, Graduate School of Agriculture, Hokkaido University, Sapporo 060-8589, Japan [[email protected]] 2 Natural History Museum, c. p. 6434, CH-1211 Geneva 6, Switzerland * Corresponding author Received 23.ii.2010, accepted ??.??.2010. Published online at www.arthropod-systematics.de on ??.??.2010. > Abstract The taxonomy, fossil record, phylogeny, and systematic placement of the booklouse family Liposcelididae (Insecta: Psocodea: ‘Psocoptera’) were reviewed. An apterous specimen from lower Eocene, erroneously identified as Embidopsocus eocenicus Nel et al., 2004 in the literature, is recognized here as an unidentified species of Liposcelis Motschulsky, 1852. It represents the oldest fossil of the genus. Phylogenetic relationships within the family presented in the recent literature were re-analyzed, based on a revised data matrix. The resulting tree was generally in agreement with that originally published, but the most basal dichotomy between the fossil taxon Cretoscelis Grimaldi & Engel, 2006 and the rest of the Liposcelididae was not supported. Monophyly of Liposcelis with respect to Troglotroctes Lienhard, 1996 is highly questionable, but the latter genus is retained because of lack of conclusive evidence. Paraphyly of Psocoptera (i.e., closer relationship between Liposcelididae and parasitic lice) is now well established, based on both morphological and molecular data. Monophyly of Phthiraptera is questionable, but support for the ‘Polyphyly of Lice Hypothesis’ is still not definitive. A checklist of valid names of all presently recognized Liposcelididae taxa (10 genera, 200 species) is also included with information on their geographical distribution. Because monophyly of the subfamily Embidopsocinae is highly questionable, we list the genera alphabetically without adopting the usual subdivision into two subfamilies. > Key words Liposcelididae, booklice, Psocoptera, Phthiraptera, parasitic lice, phylogeny. 1. Introduction The family Liposcelididae (Fig. 1) is a family of the insect order Psocodea (sensu HENNIG 1966; YOSHIZAWA & JOHNSON 2006). Within the "superorder Psocodea" (sensu HENNIG 1953), two "orders" have long been recognized, i.e., - 1 / 22 - - 1 / 22 - Psocoptera (non-parasitic members: psocids, barklice, and booklice) and Phthiraptera (parasitic members: chewing and sucking lice). However, paraphyly of Psocoptera is now widely accepted (KRISTENSEN 1991; GRIMALDI & ENGEL 2005; BESS et al. 2006). Therefore, some authors have recognized Psocodea as the only valid taxon and have rejected formal use of the order name Psocoptera (HENNIG 1966; LYAL 1985; YOSHIZAWA & JOHNSON 2006). Since LYAL (1985) proposed a close phylogenetic affinity between Liposcelididae and parasitic lice based on cladistic analysis of morphological data, the Liposcelididae are considered to be a key taxon in uncovering the origins and evolution of parasitism in lice. Liposcelididae are minute free living insects (Fig. 1) usually classified under Psocoptera, but they share a lot of features with parasitic lice (LYAL 1985; GRIMALDI & ENGEL 2005). However, the character states shared between Liposcelididae and parasitic lice are mostly reductions, and phylogenetic significance of such characters has also been questioned (LYAL 1985; YOSHIZAWA & JOHNSON 2006). Recently, several molecular-based phylogenetic analyses were performed to test Lyal's hypothesis. Results from the molecular analyses support strongly the hypothesis but, in turn, provide some novel insights into the origins and evolution of parasitism in lice. These include possibility of polyphyly of parasitic lice. Because Phthiraptera has long been recognized as one of the best supported monophyletic insect groups (HENNIG 1966; KRISTENSEN 1991; JAMIESON et al. 1999; GRIMALDI & ENGEL 2006), this result was highly surprising and is still debated. In this paper, we provide a review of the present taxonomic and systematic status of the family Liposcelididae and their relatives. This review was originally presented at the 4th Dresden Meeting on Insect Phylogeny (September 2009). The main topic at the meeting was phylogenetic importance of Liposcelididae bridging free living barklice and parasitic lice. However, taking this opportunity, we also provide more extensive review of the family including the intra-familial taxonomy and fossil records. A checklist of valid names of all currently recognized Liposcelididae taxa (10 genera, 200 species) is presented in Appendix 2. 2. Taxonomy of Liposcelididae Liposcelididae are classified under the psocodean suborder Troctomorpha. The suborder is subdivided into two infraorders, Amphientometae and Nanopsocetae. Together with Sphaeropsocidae and Pachytroctidae, Liposcelididae are assigned to the Nanopsocetae (LIENHARD & SMITHERS 2002). The parasitic lice (Phthiraptera) are close relatives of Liposcelididae (LYAL 1985; YOSHIZAWA & JOHNSON 2003; JOHNSON et al. 2004; MURREL & BARKER 2005) making Troctomorpha and Nanopsocetae both paraphyletic, unless the suborder and infraorder are re-defined to include parasitic lice. Liposcelididae are usually divided into two subfamilies, Embidopsocinae and Liposcelidinae (see below). However, the checklist in Appendix 2 does not employ this traditional system (see "Phylogeny within the family"). Except for the specialized cave- dwelling species Troglotroctes ashmoleorum (see LIENHARD 1996) all species of the Liposcelidinae have been assigned to the genus Liposcelis (ca 130 spp.). In contrast, Embidopsocinae were further subdivided into seven genera, although this subfamily contains fewer species (ca 70) than Liposcelidinae. Generally, the Embidopsocinae are considered to represent more plesiomorphic forms within the family. For example, members of Liposcelidinae are all apterous whereas - 2 / 22 - - 2 / 22 - winged forms are relatively frequent in Embidopsocinae. Monophyly of Embidopsocinae is questionable (GRIMALDI & ENGEL 2006; see also below). Genera traditionally assigned to Embidopsocinae are Belapha, Belaphopsocus, Belaphotroctes, Chaetotroctes, Embidopsocus, Embidopsocopsis and Troctulus (see LIENHARD & SMITHERS 2002). All embidopsocine genera are small, each containing less than five species, except for Belaphotroctes (19 spp.) and Embidopsocus (43 spp.). The genera Chaetotroctes, Embidopsocopsis, and Troctulus are all monotypic. The monotypic fossil genus Cretoscelis was originally considered to be the sister-group of all other Liposcelididae (GRIMALDI & ENGEL 2006; see also below). The largest genus, Liposcelis, is subdivided into four species groups (groups A, B, C, D) belonging to two sections, groups A and B to section I and groups C and D to section II. These subdivisions are based on suggestions published by BADONNEL (1962, 1963, 1967) and have more recently been defined and included in keys by LIENHARD (1990, 1998) and MOCKFORD (1993). These sections and species groups, based on usually well visible characters of tergite fusions and chaetotaxy, are very useful for organizing this large genus in practice, but their monophyly is debatable and has not yet been tested by phylogenetic analyses. Thus members of section II are characterized by a probably symplesiomorphic ‘annulate type’ of abdominal segmentation (lacking fusion of tergites), while section I is characterized by the apomorphic fusion of tergites 3–5, resulting in an abdomen of the ‘compact type’ (Fig. 1). The monotypic genus Troglotroctes is suggested by GRIMALDI & ENGEL (2006) to be imbedded phylogenetically within Liposcelis because the latter genus is, as compared to the former, characterized only by plesiomorphies. Troglotroctes is characterized by highly autapomorphic specializations related to its cave-dwelling behavior (LIENHARD 1996). Troglotroctes can be assigned to the species group D of Liposcelis on the basis of the presence of a pair of setae on the posterior half of the prosternum (see LIENHARD 1996), but this character state is probably plesiomorphic even at the level of Liposcelididae because possibly homologous setae are also present in Embidopsocinae. Therefore, monophyly of Liposcelis excluding Troglotroctes cannot be offhand rejected on the basis of available data. A key to the genera of Liposcelididae (except Cretoscelis and Troglotroctes) is given by LIENHARD (1991). LIENHARD (1990, 1998) proposes a key to the Western Palaearctic species of Liposcelis, which contains also almost all widely distributed domestic species. Some of them have a cosmopolitan distribution (see Appendix 2) and are important pests in stored food (see LIENHARD 2004b). 3. Fossil records of Liposcelididae Not many fossils are available for Liposcelididae. The oldest fossil of the family is known from the mid Cretaceous (ca 100 Mya) of Myanmar and is assigned to the monotypic genus Cretoscelis (only including C. burmitica). This genus was originally considered to represent the most basal split from the rest of the family (GRIMALDI & ENGEL 2006), but our revised data do not support this view (see below). The other known fossil species of Liposcelididae can all be assigned to extant genera (reviewed by NEL et al. 2004): Embidopsocus saxonicus (early Miocene, ca 22 Mya, see GÜNTHER 1989; upper Eocene or Miocene [?] according to NEL et al. 2004), E. eocenicus (lower Eocene,

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